Lower Cambrian Vendobionts from
China and Early Diploblast Evolution
D.-G. Shu,1,2* S. Conway Morris,3* J. Han,1Y. Li,4X.-L. Zhang,1H. Hua,1Z.-F. Zhang,1
J.-N. Liu,1J.-F. Guo,1Y. Yao,1K. Yasui5
Ediacaran assemblages immediately predate the Cambrian explosion of metazoans and should
have played a crucial role in this radiation. Their wider relationships, however, have remained
refractory and difficult to integrate with early metazoan phylogeny. Here, we describe a frondlike
fossil, Stromatoveris (S. psygmoglena sp. nov.), from the Lower Cambrian Chengjiang Lagersta ¨tte
(Yunnan, China) that is strikingly similar to Ediacaran vendobionts. The exquisite preservation
reveals closely spaced branches, probably ciliated, that appear to represent precursors of the
diagnostic comb rows of ctenophores. Therefore, this finding has important implications for the
early evolution of this phylum and related diploblasts, some of which independently evolved a
hard parts but are relatively diverse (1, 2), havea
defined community structure (3), and show a
global distribution (2). Yet their phylogenetic
interpretation remains highly controversial. Rad-
ical reassignments to lichens (4) and fungi (5)
have won little support (1, 6), but the traditional
assignments to animals (7) remain problematic.
A few taxa can be compared, with varying
degrees of reliability, to known animal groups,
including sponges (8), cnidarians (9), mollusks
(10), andarthropods(11). Most, however, remain
in phylogenetic limbo because comparisons to
either extant phyla or putative stem groups are
frustratingly imprecise. The provocative vendo-
biont hypothesis (12–14) seeks to unify disparate
taxa, including forms otherwise assigned to
groups as separate as cnidarians and arthropods,
on the basis of a distinctive body plan with a
modular quilted construction and possibly syn-
cytial tissue. If correct, this hypothesis has two
major implications. First, did at least some
vendobionts derive independently from protis-
tans, or are they a sister group of either animals
(15) or even diploblasts? Crucial in this respect
are various frondlike fossils, some of which
have been compared to the cnidarians, specifi-
cally the pennatulaceans (7, 16), whereas others
are clearly akin to other vendobionts (12, 17).
Here, we describe eight specimens of
Stromatoveris psygmoglena gen. sp. nov.,
(18) a frondose fossil from the Lower Cambrian
Chengjiang Lagerst.tte near Kunming, Yunnan,
and interpret them as a new vendobiont (Figs. 1
diacaran assemblages represent Earth_s
earliest complex macroscopic organisms
inthehistoryoflife(1). They lack skeletal
and 2). Five of them, including the holotype,
were collected near Meishucun, whereas the
other three are from Jianshan, near Haikou. Spec-
imens show preservation typical of other soft-
bodied fossils from Chengjiang and therefore
provide exceptional morphological detail. Pre-
sumably they were rapidly buried by storm
events, and most are oriented at a shallow angle
to the bedding plane. The split between part and
counterpart is therefore oblique, necessitating
composite reconstructions of each specimen
(Figs. 1, C to E, and 2, B and C).
Thebody isfoliate,withabluntly terminating
stalk that lacks obvious attachment structures
(Figs. 1; A, C, D, and F; and 2; A, B, and D).
Body length ranges between 2.5 and 7.5 cm.
Orientations were adopted for convenience and
imply no direct homologies with other orga-
nisms. Upper and lower surfaces are markedly
different. The former bears prominent branches
with quite pronounced relief. Most specimens
are too incomplete to count the precise number
but in the holotype about 15 are visible on
either side of the midline. The latter is defined
by a relatively narrow groove (Figs. 1A and 2,
A and E). The most proximal branches arise
1Early Life Institute and Key Laboratory of Continental
Dynamics, Northwest University, Xi’an 710069, China.
2School of Earth Sciences and Resources, China University of
Geosciences, Beijing 100083, China.3Department of Earth
Sciences, University of Cambridge, Downing Street, Cambridge
CB2 3EQ, UK.4College of Earth Science and Land Resources,
Chang’an University, Xi’an 710054, China.5Marine Biological
Laboratory, Graduate School of Science, Hiroshima University,
2445 Mukaishima-cho, Onomichi, Hiroshima 722-0073,
*To whom correspondence should be addressed. E-mail:
firstname.lastname@example.org (D.-G.S.); email@example.com (S.C.M.)
Fig. 1. TheCambrianvendobiontS. psygmoglena, gen. sp. nov. (A, B, and G) Holotype, ELI-Vend-05-001.
(A) Upper surface, (B) fragment (counterpart) of lower surface, and (G) enlargement of the boxed area in
(A). (C and D) ELI-Vend-05-002. (C) Composite photograph of upper and lower surfaces and (D) distal part
of upper surface removed to reveal lower surface. (E) ELI-Vend-05-003, composite photo of part and
counterpart to show both upper and lower surfaces. (F) ELI-Vend-05-004, composite photograph of part
and counterpart to show upper surface and axial rod located between upper and lower surfaces.
www.sciencemag.orgSCIENCEVOL 312 5 MAY 2006
the midline. Branches are mostly single, but oc-
casionally they bifurcate or fuse in an irregular
fashion (Figs. 1A and 2, A and E). In some
specimens, branches were filled with sediment,
suggesting that during life they were hollow
tubes. The lateral margins of the branches show
occasional short prongs or spurs. These may
represent interconnections either between the
branches or possibly into the interior of the
frond. Branch numbers appear to have shown
only a modest increase with body size, and dif-
ferences in relative width in larger specimens
grow. Distally, however, new branches can be
seen to differentiate from the surface of the
blade (Figs. 1A and 2, A and E).
The branches were evidently firmly attached
to the body, but some branches show slight im-
brication. The branches bear transverse, closely
spaced striations (Figs.1, A and G, and 2, A and
B, and fig. S1, A and D), but there is no evidence
for zooids. The regions between the branches
were narrow and recessed, and in some speci-
mensthere areassociateddark strands (Fig.1; A,
C, and E to G; and fig. S1; A, B, F, and G).
These may have been discrete structures, possibly
canal-like and presumably located in the body
may have been extensions connecting to other
regions of the body wall. Distal to the branches
the surface of the frond is smooth, although a
continuation of the midline occurs as a narrow
meandering groove (Figs. 1A and 2A). This
region was quite elongate (Figs. 1F and 2D) and
in the living animal probably acutely tapered.
Basal to the branches the midline area rapidly
widens to define a more or less smooth stalk.
The lower region broadly appears to be
divisible into two regions. More distally an ovoid
central smooth area is strongly concave(Figs. 1D
and 2B), but adjacent there are subdued ridges
the opposite direction to the branches on the up-
per surface. In addition, there is a diffuse orna-
mentation, roughly ovoid. More basally there is
subdued ribbing and a prominent leopard-skin
ornamentation, at least near the margin (Figs. 1B
and 2A). The lower surface becomes smooth
toward the basal region. One specimen (fig. S1,
C and E) shows striking arcuate structures on the
stem. Although these might be another type of
surface ornamentation, they are interpreted as
body-wall support, possibly collagenous.
(Figs. 1, C to E, and 2, C and D, and fig. S1B),
more pronounced in the stalk, suggesting its cross
section was approximately circular, whereas the
blade was somewhat compressed. As with the
branches, this sediment infill suggests the interior
across), with apparently ferruginous preservation,
is located nearer to the upper surface. In one
specimen (Figs. 1F and 2D), it has been partially
excavated, and toward the base a similar structure
occurs, but it runs transversely and possibly was
rotated to its present position as a result of decay.
The organism was benthic and embedded in
the seafloor by the stalk. Whether it lived upright
the axial structure would suggest the former
orientation, whereas the subdued morphology of
the lower surface, particularly the ovoid strongly
concavesmootharea(Figs.1D and 2B), couldbe
consistent with a recumbent mode. Mode of
feeding is conjectural and depends in part on
phylogenetic comparisons. Given the absence of
definitive zooids, one possibility is that the
particles via the narrow grooves between the
branches. Assuming a density of cilia comparable
to typical suspension feeders, this would provide a
highly effective trap. What appear to be inter-
connections between the branches (Figs. 1; A, E,
and G; and 2; A and D) suggest an exchange
system that presumably also connected to the
interior. During the organism_s life, the interior
was presumably filled with fluid or gelatinous
tissue. The central axis (Figs. 1F and 2D),
analogous to the axial rod of pennatulaceans,
presumably provided additional support, but no
other internal organs are discernible.
Although it cannot be placed in any known
genus, in overallform Stromatoveris is similar to
similar to the otherwise poorly documented
Khatyspytia (19), but the latter is more slender
and has a larger number of shorter branches.
General resemblances also exist with the fronds
Vaizitsinia (19), Charniodiscus (16, 20), Glaess-
nerina (16), and, more remotely, such forms as
Charnia (19, 20). The phylogenetic position of
these Ediacaran fronds (and by implication
Stromatoveris) is controversial (21), with oppos-
ing views favoring cnidarians, especially pen-
natulaceans (7, 16), or vendobionts (12, 14). Key
points in these differing interpretations include
attachment of the branches to the frond and
absence of unequivocal evidence for zooids, both
of which are inconsistent with the pennatulacean
hypothesis. The axial structure has an obvious
counterpart in pennatulaceans, but on functional
grounds this could be convergent. Stromatoveris
Fig. 2. Camera lucida
drawings of S. psyg-
moglena, gen. sp. nov.
(compare with Fig. 1).
The interpretative draw-
ings are composite, with
the counterpart reversed
and superimposed on the
part. Correspondences are
as follows: (A) and (E)
correspond to Fig. 1, A, B,
and G [(E) is enlargement
of boxed area in (A)]; (B),
to Fig. 1, C and D; (C), to
Fig. 1E;and (D),toFig. 1F.
5 MAY 2006VOL 312SCIENCE www.sciencemag.org
also has some similarities to the mid-Cambrian
Thaumaptilon (22), a possible Ediacaran survi-
vor. This taxon was provisionally identified as
a pennatulacean, in part on thebasis ofzooids.
The discovery of a more convincing pennatula-
cean from the Chengjiang Lagerst.tte (fig. S2)
suggests that if Thaumaptilon is a cnidarian
(assuming the zooids are correctly identified),
then it is more primitive than the anthozoans
(Fig.3). In any event, Thaumaptilon and Stromato-
veris are unlikely to be closely related.The latter
taxon lacks obvious zooids and has a markedly
different branching pattern. Branching in Stro-
matoveris also shows various irregularities
Eperhaps consistent with a less constrained
morphogenetic program (21)^ and has possible
interbranch connections, and most importantly
the distal branches differentiate from within the
upper surface. The latter arrangement is unlike
growth in pennatulaceans (or other colonial
metazoans). Such a style of growth, although
inferred in many frondose vendobionts, may
bear reexamination, especially because at least
some taxa show other distinctive character-
istics, including a striking fractal growth (17).
(and equivalent Ediacaran fossils) seems to
transcend protistan complexity. It seems likely,
therefore, that the vendobionts as currently
recognized (12, 14) are not monophyletic. Taxa
such as Ernietta and Pteridinium, built on simple
modular units and apparently with an infaunal
mode of life, may well be giant protistans (14).
The frondlike fossils, however, are interpreted as
metazoans, specifically diploblasts (Fig. 3). The
pronounced disparity within the diploblasts,
notably between cnidarians and ctenophores, has
made their early evolution highly speculative.
Dzik (23), however, has hypothesized a link
between Ediacaran fronds and Cambrian
ctenophores. Although it is difficult to accom-
modate, for instance, taxa such as Dickinsonia
and Thaumaptilon in this scheme, the fine
transverse structures seen on the branches of
Stromatoveris are similar to those seen in
Cambrian ctenophores (Fig. 3) despite their
the ciliated branches are closely spaced and
attached to the frond. In Cambrian ctenophores,
the branches became separated and the body
more globular. Both were probably benthic,
using the ciliated rows for suspension feeding,
derived. This evolutionary transition is marked
by a shift to a pelagic existence, acquisition of a
gelatinous body plan, and co-opting of the ciliary
rows from feeding to locomotion.
Stromatoveris joins a select group of Edi-
acaransurvivors(22–25). In comparison to those
Cambrian survivors showing typical Ediacaran-
like preservation (24, 25), the material shows
new features unobservable in the coarser host
matrix. Quality of preservation matches that of
Thaumaptilon (22), but as noted the similarities
between the two taxa are evidently convergent
(Fig. 3). The possible example of an Ediacaran
survivor, reported earlier from Chengjiang (26),
has no similarity to the example described here,
and its wider relationships are uncertain.
References and Notes
1. G. M. Narbonne, Annu. Rev. Earth Planet. Sci. 33, 421
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Study (Cambridge Univ. Press, Cambridge, 1984).
8. J. G. Gehling, J. K. Rigby, J. Paleontol. 70, 185 (1996).
9. A. Y. Ivantsov, M. A. Fedonkin, Palaeontology 45, 1219
10. M. A. Fedonkin, B. M. Waggoner, Nature 388, 868 (1997).
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Palaeontology 46, 447 (2003).
12. A. Seilacher, J. Geol. Soc. London 149, 607 (1992).
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7, 43 (2003).
15. L. W. Buss, A. Seilacher, Paleobiology 20, 1 (1994).
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17. G. M. Narbonne, Science 305, 1141 (2004); published
online 15 July 2004 (10.1126/science.1099727).
18. Systematic paleontology is as follows: Stem group
Ctenophora (in part Vendobionta). Order ‘‘Charniomorpha’’
(27). Family Stromatoveridae Shu, Conway Morris,
and Han fam. nov. genus Stromatoveris Shu, Conway
Morris, and Han gen. nov. Stromatoveris psygmoglena
Shu, Conway Morris, and Han sp. nov. Etymology: Generic
name is compound, based on mattress (Greek stromatos)
and Spring (Latin veris), oblique references to the
mattresslike fossil and its discovery near the city of
Kunming, nicknamed Spring City; specific name implies
wonderful (Greek glenos) fan (Greek psygma). Holotype:
ELI-Vend-05-001. Other material: ELI-Vend-05-002 to
008. Stratigraphy and locality: Heilinpu (formerly
Qiongzhusi) Formation, Yu’anshan Member (Eoredlichia
zone), Lower Cambrian. Specimens collected in
Meishucun and Jianshan areas. Diagnosis: Foliate, divided
into basal stalk with rounded termination and flatter frond
that tapers distally. Body strongly differentiated, both
between upper and lower surfaces and also along axis of
both upper and lower surfaces. Upper surface with
prominent attached branches in midregion, strongly acute
insertion, separated by central groove, branches differen-
tiate from surface in distal region. Branches bear fine
transverse structures. Basal and distal regions of upper
surface smooth. Distally lower surface has a medial
elliptical smooth area from which radiate lines with ovoid
ornamentation. Basal region bears subdued ribbing and
prominent oval ornamentation, but toward basal tip
19. M. A. Fedonkin, in The Vendian System, B. S. Sokolov,
A. B. Iwanowski, Eds. (Springer-Verlag, Berlin, 1990),
vol. 1, pp. 71–120.
Fig. 3. Outline of metazoan phylogeny, showing proposed position of Stromatoveris and Thaumaptilon
(22) as primitive ctenophores and cnidarians, respectively, so implying convergent evolution of a frondlike
habit. Metazoan phylogeny is still in a state of flux, but here sponges are taken to be basal, with the
calcareans possibly a sister group of all other metazoans (28). The position of the placozoans is
controversial, but here they are treated as primitive diploblasts (29), evolving before the invention of
nerve cells (30). Ctenophores are monophyletic (31) and are taken to be the sister group of cnidarians
plus triploblasts (28). As argued in the text, ctenophores were primitively frondlike (vendobionts) before
acquiring a globular body with separate comb rows that eventually were used in a pelagic existence.
Although ctenophores have a biradial symmetry, this has a unique rotational element and may be derived
and effectively unrelated to the biradial symmetry that may be primitive to cnidarians. Cnidarians are also
monophyletic and are divided into anthozoans and medusozoans (32). Although previously ctenophores
have been argued to be the sister group of all bilaterians, it is now widely accepted that cnidarians are the
sister group (33). The triploblasts are composed of deuterostomes and protostomes.
www.sciencemag.orgSCIENCE VOL 312 5 MAY 2006
20. T. D. Ford, Proc. Yorks. Geol. Soc. 31, 211 (1958). Download full-text
21. B. N. Runnegar, Neues Jahrb. Geol. Pala ¨ontol. Abh. 195,
22. S. Conway Morris, Palaeontology 36, 593 (1993).
23. J. Dzik, J. Morphol. 252, 315 (2002).
24. S. Jensen, J. G. Gehling, M. L. Droser, Nature 393, 567
25. J. W. Hagadorn, C. M. Fedo, B. M. Waggoner, J. Paleontol.
74, 731 (2000).
26. W.-T. Zhang, L. Babcock, Acta Palaeontol. Sinica 40
(Suppl.), 201 (2001).
27. The phylogenetic questions associated with Ediacaran taxa
have largely precluded a higher-level taxonomy.This ordinal
designation would include both the taxon described here
andsuchgeneraasCharniodiscus, Glaessnerina, Khatyspytia,
Vaizitsinia, and possibly Charnia.
28. M. Medina, A. G. Collins, J. D. Silberman, M. L. Sogin,
Proc. Natl. Acad. Sci. U.S.A. 98, 9707 (2001).
29. A. Ender, B. Schierwater, Mol. Biol. Evol. 20, 130 (2003).
30. T. Hadrys, R. DeSalle, S. Sagasser, N. Fischer, B. Schierwater,
Mol. Biol. Evol. 22, 1569 (2005).
31. M. Podar, S. H. D. Haddock, M. L. Sogin, G. R. Harbison,
Mol. Phyl. Evol. 21, 218 (2001).
32. A. G. Collins et al., Syst. Biol. 55, 97 (2006).
33. K. J. Peterson, M. McPeek, D. A. D. Evans, Paleobiology 2
(Suppl.), 36 (2005).
34. We thank Y. Ji, M. Cheng, and J. Zhai at the Early Life
Institute; S. Last, S. Capon, and V. Brown at Cambridge
for technical assistance; and N. Butterfield and three
anonymous referees for critical remarks. The work is
supported by the National Natural Science Foundation of
China, Ministry of Science and Technology of China,
Program for Changjiang Scholars and Innovative Research
Team in University (PCSIRT), the Cowper-Reed Fund, and
St. John’s College, Cambridge.
Supporting Online Material
Figs. S1 and S2
4 January 2006; accepted 31 March 2006
Defective Lipolysis and Altered Energy
Metabolism in Mice Lacking Adipose
Guenter Haemmerle,1Achim Lass,1Robert Zimmermann,1Gregor Gorkiewicz,2Carola Meyer,5
Jan Rozman,5Gerhard Heldmaier,5Robert Maier,3Christian Theussl,6Sandra Eder,1
Dagmar Kratky,4Erwin F. Wagner,6Martin Klingenspor,5Gerald Hoefler,2Rudolf Zechner1*
Fat tissue is the most important energy depot in vertebrates. The release of free fatty acids (FFAs)
from stored fat requires the enzymatic activity of lipases. We showed that genetic inactivation of
adipose triglyceride lipase (ATGL) in mice increases adipose mass and leads to triacylglycerol
deposition in multiple tissues. ATGL-deficient mice accumulated large amounts of lipid in the heart,
causing cardiac dysfunction and premature death. Defective cold adaptation indicated that the
enzyme provides FFAs to fuel thermogenesis. The reduced availability of ATGL-derived FFAs leads to
increased glucose use, increased glucose tolerance, and increased insulin sensitivity. These results
indicate that ATGL is rate limiting in the catabolism of cellular fat depots and plays an important
role in energy homeostasis.
olism. Disruptions of this balance underlie
metabolic diseases such as obesity and type
II diabetes (1–3). Hormone-sensitive lipase
(HSL) was once thought to be the major
enzyme responsible for the lipolytic break-
down of cellular fat stores (4–6). However,
HSL-deficient mice are lean, and they efficient-
ly mobilize FFAs from triacylglycerol (TG)
stores (7, 8), suggesting that other TG hydro-
lases play an important role. Recently, we and
others reported the discovery of an enzyme
that we named in accordance with its phys-
iological activity: adipose triglyceride lipase
(ATGL) (9–11). Other designations for this
enzyme have been: transport secretion protein
(TTS), desnutrin (10), calcium-independent
dipose tissue mass in mammals is
determined by the dynamic equilibri-
um of lipid synthesis and lipid catab-
phospholipase A2z (iPL-A2z) (11), adiposome
triglyceride lipase (ATGL) (12), and patatin-like
phospholipase domain–containing protein 2
(PNPLP2). ATGL specifically hydrolyses
long-chain fatty acid TG (9, 11) and is pre-
dominantly expressed in adipose tissue and,
to a lesser extent, in cardiac muscle, skeletal
muscle, testis tissue, and other tissues. The
finding that ATGL mRNA expression is
regulated by fasting/feeding (10) as well as
hormones and cytokines (13, 14), and that the
inhibition of ATGL in vitro (9, 12) markedly
decreases TG catabolism, imply that the en-
zyme plays an important role in lipolysis.
To elucidate the physiological function of
ATGL during lipid mobilization in vivo, we
inactivated the Atgl gene in mice by replacing
the first exon, including the translational start
codon and the lipase consensus sequence motif
(GXSXG, where G is Gly, S is Ser, and X is
any amino acid), with a neomycin expression
cassette (fig. S1). Atgl(j/j) mice showed
accumulation of neutral lipid to supraphysio-
logical levels in most tissues, suggesting an
essential role for ATGL in cellular TG
catabolism (table S1). The TG content in tissues
of Atgl(þ/j) mice resembled that of wild-type
(WT) mice, except for cardiac muscle where
there was a twofold increase. Compared with
WT mice, Atgl(j/j) animals were heavier
(Fig. 1A and fig. S2A), displayed a twofold in-
crease in whole body fat mass, and exhibited
enlarged adipose fat depots (Fig. 1B and fig.
S2, B and C). The mutants had an increased
wet weight of gonadal (2.1-fold) and inguinal
white adipose tissue (WAT) (1.6-fold), as well
as intrascapular brown adipose tissue (BAT)
(8.5-fold). Ad libitum food intake E3.6 T 0.6 g
in Atgl(j/j) mice and 3.3 T 0.7 g in WT mice^
and lean body mass (Fig. 1A) were similar in
mice of both genotypes. Consistent with in-
creased adipose weight, plasma leptin levels
were elevated in fed (2.1-fold) and fasted (4.4-
fold) Atgl(j/j) mice (table S2). The leptin/fat
mass ratio, however, was similar in mutant and
control mice. Morphological analyses of adipose
tissue from Atgl(j/j) mice (fig. S2C) revealed
enlarged lipid droplets in white E4690 T 235
mm2for Atgl(j/j) versus 3382 T 90 mm2for
WT^ and brown adipocytes E1395 T 119 mm2
for Atgl(j/j) versus 67 T 10 mm2for WT; n 0
100 cells analyzed, P G 0.001^. The multilocular
lipid droplets normally observed in BAT became
unilocular and strongly resembled white fat in
appearance. Thus, the absence of ATGL in mice
causes fat cell hypertrophy and mild obesity.
In cardiac muscle, ATGL deficiency caused
severe TG accumulation (Fig. 1, C and E). At
the age of 12 weeks, mice had a TG content
in myocytes more than 20 times higher in
Atgl(j/j) mice than in WT controls, causing a
1.4-fold increase in heart weight E193 T 18 mg
for Atgl(j/j) versus 131 T 12 mg for WT, P G
0.001, n 0 8^. Histological analyses revealed an
age-dependent increase of lipid droplets in
number and size in cardiomyocytes, starting
with microvesicular lipid accumulation at the
age of 6 weeks and progressing to the accu-
mulation of large droplets at the age of 18
weeks (Fig. 1E). This massive lipid buildup led
to severe cardiac insufficiency. In echocardiog-
raphy (Fig. 1E and table S3), the ejection
fraction of the left ventricle was drastically
reduced in Atgl(j/j) mice (40.2 T 26.5%)
compared with WT (80.5 T 17.1%, P G 0.001).
Additionally, a marked disturbance of cardiac
texture and increased fibrosis was noted (Fig.
1E). The interventricular septum (1.9 T 0.6 mm)
1Institute of Molecular Biosciences, University of Graz,
Center of Molecular Medicine, Medical University of Graz,
Biology, Philipps-University Marburg, Germany.6Research
Institute of Molecular Pathology, Vienna, Austria.
2Institute of Pathology;
4Institute of Molecular Biology and Biochemistry,
3Department of Cardiol-
5Department of Animal Physiology, Faculty of
*To whom correspondence should be addressed. E-mail:
5 MAY 2006VOL 312SCIENCEwww.sciencemag.org